d N1 N N

[ ]
3. Nondimensionalize
{ dt
=r 1 N 1 1− 1 −b 12 2

d N2
dt
K1

[ N
K1

=r 2 N 2 1−b21 1
K2 ]
N1 N2 r2 K2 K1
Let u1= , u2= , τ =r 1 t , ρ= , a 12=b12 , a 21=b21
K1 K2 r1 K1 K2

K 1 d u1 u K K a u K du
[ ]
{
=r 1 u1 K 1 1− 1 1 − 1 12 2 2 ❑ 1 =u1 [ 1−u1−a12 u2 ]
dτ K1 K 2 K 1 ⇒ dτ
r1
K 2 d u2 K a u K du
dτ
r1
[ K 1 K 2 ⇒ dτ ]
=r 2 u 2 K 2 1− 2 21 1 1 ❑ 2 = ρu2 [ 1−a 21 u1 ]

Steady States and phase plane singularities:

d u 1¿ d u2¿
= =0, there are three equilibrium/critical points:
dt dt
¿ ¿
 u1 =0∧u2 =0
¿ ¿
 u1 =1∧u2 =0
1
−1
 ¿ 1 ¿ a 21 1−a21
u1 = ∧u 2 = = ,a ≤1
a21 a 12 a12 a21 21

Stability and Phase Plane Trajectories:

1−2 u1−a12 u2 −a12 u 1
Jacobian matrix A=
[
−ρ u2 a21 ρ ( 1−a21 u 1 ) ]
¿ ¿
At u1 =0∧u2 =0 ,| A−λI |=
0
ρ−λ [ 1−λ0 ]
=0❑ (1−λ ) ( ρ− λ )=0
⇒

❑ λ 1=1 , λ2=ρ which are both positive, hence steady state is unstable.
⇒
−1−λ −a12
¿ ¿
At u1 =1∧u2 =0 ,| A−λI|=
0 [
ρ ( 1−a 21) −λ
=0
]
❑ (−1−λ ) ( ρ ( 1−a21 )−λ )=0❑ λ 1=−1 , λ2=ρ ( 1−a21 ), 1−a21should be less
⇒ ⇒
stable if a21>1
than zero to make the steady state stable. {
unstable if a21 <1
¿ 1 ¿ 1−a21
For u1 = ∧u 2 = , a ≤ 1it is easier to use phase plane in analysis. Since its
a21 a12 a21 21
2 a21−3 2 a21−3 2 ρ ( 1−a21 )
eigenvalues are quite complicated:
λ 1 , λ1=
a21
±
√( a21
2
) +4
a21 .
 1−u 1−a12 u 2=0❑ intercepts : 0 ,
⇒ ( a1 )∧(1 , 0)
12
1
1−a21 u1=0❑ u1 =
⇒ a 21

Consider a 21>1:
For Area 1:
1−u1
1−u 1−a12 u 2> 0❑ u2 <
⇒ a 12
1
1−a21 u1 >0❑ u1 <
⇒ a21
1−u 1 d u 1 0∧d u2
Thus, u1 +a 12 u2 <u1 +a12 =1. Hence, > > 0.
a 12 dτ dτ
For Area 2:
1−u1
1−u 1−a12 u 2< 0❑ u2 >
⇒ a 12
1
1−a21 u1 >0❑ u1 <
⇒ a21
1−u 1 d u 1 0∧d u2
Thus, u1 +a 12 u2 >u1 +a12 =1. Hence, < > 0.
a 12 dτ dτ
For Area 3:
1−u1
1−u 1−a12 u 2> 0❑ u2 <
⇒ a 12
1
1−a21 u1 <0❑ u1 >
⇒a21
d u 1 0∧d u2
Hence, > < 0.
dτ dτ
For Area 4:
1−u1
1−u 1−a12 u 2< 0❑ u2 >
⇒ a 12
1
1−a21 u1 <0❑ u1 >
⇒ a21
d u 1 0∧d u2
Hence, < < 0.
dτ dτ

U2

Area 2
1 1−u 1−a12 u 2=0
Area 4 separatrix
a12

1 1−a21
Area 1 ( ,
a21 a12 a 21 )